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Maternal Care in Omaspides Bistriata Boheman (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini)

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Maternal Care in Omaspides Bistriata Boheman (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) www.biotaxa.org/rce. ISSN 0718-8994 (online) Revista Chilena de Entomología (2020) 46 (4): 613-622. Research Article Maternal care in Omaspides bistriata Boheman (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) Cuidado maternal en Omaspides bistriata Boheman (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) Rolando Ramírez C.1 and Marcela Sánchez-Ocampo2 1Natural and Exact Sciences School (ECEN), Natural resource management (MARENA), Universidad Estatal a Distancia (UNED), San José, Costa Rica. [email protected] 2 National Museum, Natural History Department, San José, Costa Rica. [email protected] ZooBank: urn:lsid:zoobank.org:pub: EEA83797-4FCE-4A94-BB6F-CDE31400A1DC https://doi.org/10.35249/rche.46.4.20.07 Abstract. Maternal care (subsociality): characterization of the different stages of maternal care and its efficiency as a strategy. Maternal care and larval development of Omaspides bistriata Boheman, 1862 (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini) are described; including characteristics and manner in which maternal care is given across the different stages of development of the specie (eggs, larvae, pupae and teneral adults). We report the oviposition of eggs, the duration to hatch the eggs, and the duration of larval period, pupal stage, and emergence. A life table and survival curve is presented covering all life stages. Changes in the behavior and feeding habits are also noted for the immatures and the attending mother. Key words: Beetle, behavior, hostplants, parental care. Resumen. Cuidado maternal (subsocialidad): caracterización de las diferentes etapas del cuidado materno y su eficiencia como estrategia. Se describen el cuidado maternal y el desarrollo larvario de Omaspides bistriata Boheman, 1862 (Coleoptera: Chrysomelidae: Cassidinae: Mesomphaliini); incluyendo características y forma en que se brinda el cuidado materno en las diferentes etapas de desarrollo de la especie (huevos, larvas, pupas y adultos tenerales). Informamos la oviposición de los huevos, la duración de la eclosión de los huevos y la duración del periodo larvario, estado de pupa y emergencia. Se presenta una tabla de vida y una curva de supervivencia que cubre todas las etapas de la vida. También se notan cambios en el comportamiento y en los hábitos alimentarios de los inmaduros y de la madre que los atiende. Palabras clave: Comportamiento, cuidado maternal, escarabajo, planta hospedera. Introduction Subsociality (i.e. extended parental care) is the most widespread social behavior among insects (Royle et al. 2012). It is known to occur in 167 families of 19 insect orders (Machado & Trumbo 2018). Subsocial behavior encompasses care for offspring by protection against natural enemies, guiding towards or provisioning of food resources during most or all Received 23 September 2020 / Accepted 22 October 2020 / Published online 30 October 2020 Responsible Editor: José Mondaca E. Este es un artículo de acceso abierto distribuido bajo los términos de la licencia Creative Commons License (CC BY NC 4.0) Ramírez and Sánchez-Ocampo: Maternal care in Omaspides bistriata Boheman. immature stages (Clutton-Brock 1991; Royle et al. 2012). Within Coleoptera, subsociality is a rare phenomenon and the behavior has been recorded in 12-17 out of 176 beetle families (Chaboo et al. 2014; Leocádio et al. 2020). Chrysomelidae Latreille, 1802 is one of the beetle families in which this behavior has evolved indepentently in two subfamilies of 15: Cassidinae Gyllenhal, 1813 and Chrysomelinae Gyllenhal, 1802 (Windsor & Choe 1994; Chaboo et al. 2014; Leocádio et al. 2020). It is noteworthy that many Chrysomelidae larvae feed on the external tissues of plants (exophytic) (White 1983; Jolivet 1997), being more exposed to predation or parasitism than larvae of most Coleoptera, which live and hide in logs or underground (Wilson 1971; White 1983; Costa 2006). Many exophytic insects developed parental care, probably in a response to their feeding behavior (Cornell & Hawkins 1995). Cassidinae sensu lato is the second largest subfamily of leaf beetles with 6,300 species, distributed worldwide (Borowiec & Świętojańska 2018). Within Cassidinae, parental care has evolved independently in two closely related lineages that feed on distinct host plants families: Asteraceae and Convolvulaceae (Leocádio et al. 2020). Until now, parental has been recorded in 24 species belonging to five genera (Chabooet al. 2014; Flinte et al. 2015; Macedo et al. 2015). The genus Omaspides Chevrolat, 1836 (Mesomphaliini) belongs to the Convolvulaceae feeding clade (along with Acromis Chevrolat, 1836 and Paraselenis Spaeth, 1913) and parental care has been recorded in 11 out of its 40 recognized species (Gomes et al. 2012; Chaboo et al. 2014). Parental care and number of eggs per clutcth were briefly described for Omaspides (Omaspides) bistriata Boheman in Panama (Windsor & Choe 1994). This genus has three subgenera with 40 recognized species. Here we present a detailed life history description for O. (O.) bistriata in Costa Rica, including development time of all imature stages and number of individuals per clutch with survival rates for each stage. Materials and Methods Study area The Veragua Rainforest Park is located at the foothills of Fila Matama sector, in La Amistad Caribe International Park (Costa Rica-Panamá), Limón, Costa Rica within the very humid tropical forest life zone (bmh-T) (Holdridge 1967). The monitoring and specimen collection were carried out within an area composed mainly of secondary forest at an average elevation of 430 m (N 9°55’35.7’’ - W 83°11’27.9’’). With a minimum average annual temperature of 19.6 °C and a maximum of 27.1 °C. The vegetation surrounding the Veragua Rainforest reserve was monitored between December (2008) and February (2011), observing the behavior and development of individuals of the species O. (O.) bistriata. In the monitored area, there are high altitude trees distant from each other, so the canopy is not complete, and this enables the entrance of light and the formation of clearings which are, in turn, favorable for the proliferation of herbaceous, shrubs and vines, among which is the morning-glory Ipomoea purpurea (L.) Roth (Convolvulaceae), the host plant of O. (O.) bistriata. Biological studies Three couples in copula were spotted on three distinct occasions on February 4, 2009; December 27, 2009 and January 16, 2011. In order to monitor their reproductive behavior, the sites were marked and visited daily for at least 6 hours a day. Besides the three caring females and offspring, we found four eggs clutches without 614 Revista Chilena de Entomología 46 (4) 2020 parental attendence and they were also monitored. An additional female caring for pupal offspring was monitored. The females remain feeding and sleeping in the plants where the copulation occurs. After three to four days it was observed that the volume of the females’ abdomen increased gradually, until the time of eggs laying. The oviposition of the first monitored female was performed on February 27, 2009 and was composed of 32 eggs, while the second female was oviposited on January 16, 2010 with 34 eggs and the third on January 30, 2011 with 29 eggs. All egg layings were monitored during their development and care process, up to adulthood individuals. In addition, four eggs clusters were found (January 2009 with 28 eggs, March 2009 with 35 eggs, April 2010 with 30 eggs and January 2011 with 31 eggs) with no adult individuals present and in January 2011 we found a mother caring for seven pupae. Data analyss Based on the analysis and on the data of the three eggs layings and their complete development cycles, a life table was created with their respective survival curve, the percentage of viable adults, completing development the average number of oviopositions, survival to larval and pupal period, hatching of juvenile individuals and sexual adults at the end of the cycle observed under natural conditions. To determine the values of the life table an average of the data between the three ovipositions was taken and classified of the following form: • X = Age in days, starting from eggs laying • nx = Average number of living individuals at age X in days • lx = Proportion of surviving individuals at the beginning of the age range X • dx = Number of individuals killed during the age range X to X +1 • qx = Per capita mortality rate during the interval X to X +1 Where: • lx = nx / n0 nX+1 = nx – dx qx = dx / nx To plot the exponential survival curve data were used: • Axis X = Age in days, from oviposition (X) • Axis Y = Average number of living individuals at age X Specimen Voucher All samples were deposited at Veragua Rain Forest biological collection. Host plant was determinated by Fabian Araya (Universidad Nacional) and beetles were deteminated by Rolando Ramírez (Universidad Estatal a Distancia). Results Live cycle The complete life cycle lasts 44-50 days as can be seen in Table 1. Eggs. Oviposition occurred 19 days after copulation. The eggs measured 1-1.5 mm long, and they hatched after 5-6 days of incubation. The eggs were light orange in color, and in clutches of 29 to 34 eggs on abaxial surface of host plant leaves (Figs. 1A, 1B). Larvae. Larval stages develop in 34 days average, with a length from 1.5-2 mm on the 615 Ramírez and Sánchez-Ocampo: Maternal care in Omaspides bistriata Boheman. first instar to 9-10 mm on the last one, highlighting the formation of the circular exuvial- fecal shield (cycloalexy), with the head inwards in the group of larvae during their early stages. In the first stages, the larvae fed mainly on mature leaves, while in the later stages they ate tender, mature leaves and even the stem. When the larvae move through the plant, the mother travels the entire area in which they are found, and at night they regroup again. (Figs. 1C, 1D). Pupae. Between days 38 and 42. The pupae began to form (Figs. 1E, 1F), presenting a size between 8-10 mm, they were attached in both the bundle and the underside of the leaves, mostly individually, however, some were observed in groups of two to four pupae.
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